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Supplementary Studies on Ant 

Larvae: Cerapachyinae, 

Pseudomyrmecinae and 

Myrmi cinae 



BY GEORGE C. WHEELER 

and 
JEANETTE WHEELER 



Reprinted from PSYCHE, Vol. 80, No. 3, September 1973 
pages 204-211 



W1UJ6M U BROWN 



-ill? 



SUPPLEMENTARY STUDIES ON ANT LARVAE: 

CERAPACHYINAE, PSEUDOMYRMECINAE 

AND MYRMICINAE* 

BY GEORGE C. WHEELER AND JEANETTE WHEELER 

Laboratory of Desert Biology, Desert Research Institute, 

University of Nevada System, Reno 89507 

Subsequent to the publication of our first supplement on the ant 
larvae of the subfamily Cerapachyinae (1964), our first paper on 
Pseudomyrmecinae (1956) and several supplements on Myrmicinae 
(1960, 1972, 1973) we have received from other myrmecologists so 
much additional material that it has now become necessary to pub- 
lish a supplement. 

Genus PHYRACACES Emery 

REVISION : The last sentence of our generic characterization 
(1964: 69) should read: Hypopharynx usually spinulose dorsally. 

Phyracaces elegans Wheeler (Fig. 2). Length (through spiracles) 
about 4.7 mm. Very similar to Ph. larvatus (1964: 69) except as 
follows. Body more slender. A pair of bosses on lateral surfaces of 
venter of AI-AVI. Spiracles small, AI largest, diameter decreasing 
posteriorly. Integument densely spinulose, spinules in short to long, 
subtransverse to arcuate rows. Body hairs less numerous and shorter 
(0.025-0.05 mm long). Head hairs shorter (0.009-0.019 mm long). 
Posterior surface of labrum with a ventrally directed medial boss 
bearing 6 sensilla, about 5 sensilla on each lateral surface. Mandi- 
bles with narrower base. No spinules seen on hypopharynx. 

YOUNG LARVA: Length (through spiracles) about 2.1 mm. Simi- 
lar to mature larva above except as follows. Neck curved, abdomen 
with straight ventral profile and C-shaped dorsal profile. Body hairs 
shorter (0.01-0.033 mm long). Integument spinulose, spinules mi- 
nute, isolated laterally and in short rows dorsally and ventrally. 
Antennae less distinct. Maxillae lacking spinules; galea a slightly 
raised pair of sensilla. No spinules seen on labium ; opening of 
sericteries a short slit. 

VERY YOUNG LARVA: Length (through spiracles) about 1.5 mm. 
Entire body arcuate ventrally. Otherwise similar to young larva. 

Material studied: numerous larvae from New South Wales, 
courtesy of Rev. B. B. Lowery. 

*Manuscript received by the editor September 28, 19/3 

2O4 



1973] 



Wheeler sf Wheeler Ant Larvae 



205 




Fig. 1. Cerapachys (Syria) australis: a, head in anterior view, X95; 
b, left mandible in anterior view, X314; c, larva in side view, X22; 
d and e, two types of body hairs, X444. Fig. 2. Phyracaces elegans: head 
in anterior view, X 74. Fig. 3. Tetraponera natalensis: left antenna in 
lateral view, X339. 



Phyracaces ficosus Wheeler. Length (through spiracles) about 
4.4 mm. Very similar to Ph. larvatus (1964: 69) except in the 
following details. Spiracles on first abdominal somite slightly larger, 
remainder small and subequal. Body hairs shorter (0.013-0.063 mm 
long). Antennae with 2 sensilla each. Head hairs shorter (0.008- 
0.025 mm long). Galeae digitiform. (Material studied: 14 larvae 
from New South Wales, courtesy of Rev. B. B. Lowery.) 

Genus CERAPACHYS F. Smith 

REVISION: Our generic characterization (1964: 67) should be 
replaced with the following : Leg vestiges small paraboloidal papillae. 
Body hairs usually simple. Head (including mouth parts) subpyri- 
form in anterior view. Head hairs usually short. Mandibles long, 
slender and with median border erose. Maxillary palp short; galea 
long and digitiform. 

Cerapachys (Syscia) australis Forel (Fig. i). Length (through 
spiracles) about 3.2 mm. Body long and subcylindrical ; about 12 



206 Psyche [September 

differentiated somites; head on anterior end; a small posteriorly 
projecting boss on AX. Anus ventral. Spiracles small. Entire 
integument densely spinulose, spinules minute and in short to long 
straight or arcuate rows. Body hairs short, uniformly distributed 
and moderately numerous. Of two types: (i) 0.025-0.063 mm long, 
mostly bifid, sometimes with one or both branches rebranched, on all 
somites; (2) 0.037-0.05 mm long, simple, a few on each somite. 
Cranium subhexagonal in anterior view, slightly longer than wide. 
Antennae large, each a low mound with 3 minute sensilla, each 
bearing a minute spinule. Head hairs few, 0.025-0.05 mm long, 
simple or bifid. Labrum subarcuate, about twice as wide as long; 
anterior surface with 8 sensilla on and near ventral border; posterior 
surface with about 6 sensilla ventromedially and with a few oblique 
arcuate rows of minute spinules. Mandibles narrowly subtriangular 
in anterior view; apex rather long, narrow and heavily sclerotized; 
medial border with 6-8 small denticles. Maxillae with apex para- 
boloidal and sparsely spinulose, spinules minute to short and in a 
few arcuate rows; palp a peg with 4 (2 encapsulated and 2 bearing 
a spinule each) apical and one lateral sensilla; galea digitiform with 
2 apical sensilla, each bearing a minute spinule. Labium subtrape- 
zoidal, widest distally, anterior surface densely spinulose, spinules 
minute and in numerous short arcuate rows ; palp a rounded elevation 
with 5 (2 encapsulated and 3 bearing a spinule each) sensilla; an 
isolated sensillum between each palp and opening of sericteries; the 
latter a slit in a shallow depression on anterior surface. Hypopharynx 
with minute spinules in long transverse sub-parallel rows. (Material 
studied: 10 larvae from Queensland, courtesy of Rev. B. B. Lowery.) 

SUBFAMILY PSEUDOMYRMECINAE 

We have never been able to key the genera of this subfamily. 
Except for head shape, where the difference in the species of Pachy- 
sima is greater than that between any two genera, some of the vari- 
ants of any character in any genus can be found in other genera. 

Bernard (1951 : 1053) included larval characters in his character- 
ization of the subfamily, which he called family Promyrmicidae. 

Sudd (1967: 123) discussed the feeding of the larvae. He stated 
(erroneously) that the trophothylax was formed by the bases of the 
'rudimentary legs; we have shown (1956: 375, 383) that it is 
"formed from the depressed ventral surface of the thorax and elab- 
oration of the first and second abdominal somites." 



1973] W 'heeler & Wheeler Ant Larvae 2OJ 

Genus PSEUDOMYRMEX Lund 

Janzen (1967: 344). Beltian bodies are cut up by the workers 
and fed to the larvae. 

The following species of Pseudiomyrmex are compared with Ps. 
alliodtorae 1956: 379); only differences are given here. 

Pseudomyrmex adustus Borgmeier. Length (through spiracles) 
about 4.8 mm; straight length about 4.6 mm. Body hairs: (2) 
0.05-0.25 mm long, longest of AI-AV; (3) 0.2-0.25 mm long, 2 
only on each Ti-3 and AI-AIII. Head hairs more numerous and 
0.05-0.25 mm long. Posterior surface of labrum with a cluster of 3 
sensilla in the middle of each half. ( Material studied : 7 larvae from 
Brazil, courtesy of Dr. K. Lenko.) 

Pseudomyrmex belti fulvescens Emery (= Ps. ferrugineus F. 
Smith). Janzen 1967: Description p. 394; feeding of larvae p. 416- 
417; handling of larvae p. 418. Similar information in Janzen 
1966. 

Pseudomyrmex elongatus(Borgmeier\^L&ngth (through spiracles) 
about 3.8 mm; straight lengHT" aBbut 3.6 mm. Body hairs longer: 
(i) 0.006-0.018 mm long; (2) 0.018-0.2 mm long; (3) 0.175-0.22 
mm long, 4 in a row across the dorsum of each Ti-3 and AI-AIV. 
Head hairs more numerous and slightly longer (0.01-0.05 mm long). 
(Material studied: 15 larvae from Brazil, courtesy of Dr. K. Lenko.) 

Pseudomyrmex schuppi Forel. Length (through spiracles) about 
5.9 mm; straight length about 5.7 mm. Largest spiracles on AI. 
Body hairs longer: (i) 0.013-0.025 mm long; (2) 0.038-0.25 mm 
long; (3) 0.25-0.33 mm long, 4 in a row across the dorsum of each 
Ti-3 and AI-AIII. (Material studied: numerous larvae from Brazil, 
courtesy of Dr. K. Lenko.) 

Pseudomyrmex subtilissimus Emery. Length (through spiracles) 
about 4.3 mm; straight length about 3.9 mm. Body hairs (i) 0.006- 
0.018 mm long; (2) 0.018-0.15 mm long; (3) about 0.15 mm long, 
4 in a row across the dorsum of each Ti-3 and AI-AIV. Head hairs 
slightly longer (0.025-0.05 mm long). (Material studied: 6 larvae 
from Brazil, courtesy of Dr. K. Lenko.) 

Pseudomyrmex termitarius F. Smith. Length (through spiracles) 
about 5.1 mm; straight length about 4.7 mm. Body stouter. Body 
hairs about twice as numerous: (i) 0.013-0.05 mm long; (2) 0.05- 
0.275 mm long; (3) 0.25-0.35 mm long, 4 in a row across dorsum 
of each Ti-3 and AI-AIV. Head hairs more numerous, longer 
(0.013-0.05 mm long) and finely denticulate. Labrum with width 
twice the length, with anterior lobes more prominent and with 2 



2O8 Psyche [September 

minute hairs on anterior surface. Mandibles with teeth stouter and 
blunter; lateral outline less curved; denticles on anterior surface 
more numerous. Maxillary apex less constricted and with spinules 
longer and covering a greater portion of the surface. Labium with 
more numerous spinules. (Material studied: 9 larvae from Brazil, 
courtesy of Dr. K. Lenko.) 

Genus TETRAPONERA F. Smith 

Tetraponera natalensis F. Smith (Fig. 3). Length (through spira- 
cles) about 8.2 mm; straight length about 6.2 mm. Similar to T. 
aitkeni (1956: 388) except as follows. Body slightly stouter at AV 
and AVI. Integument of AIX and AX with minute spinules. Body 
hairs: (i) 0.008-0.075 mm long; (2) 0.025-0.15 mm long, longest 
with tip branched or denticulate; (3) 0.175-0.3 mm long, 4 in a 
row across the dorsum of each Ti-3 and AI-AVI. Each antenna 
represented by 3 individually raised sensilla on a small base. Head 
hairs longer (0.0130.11 mm long) and less numerous, with or 
without alveolus and articular membrane, some with denticles near 
the tip. Labrum with breadth less than twice length; borders sinu- 
ate; anterior surface with 6 sensilla and 2 hairs on each half; pos- 
terior surface with 9 sensilla on each half; spinules as in T. aitkeni. 
Anteromedial surface of mandibles with large spinules, which are 
isolated or in short rows of 2 or 3. Maxillae with rather numerous 
long spinules in short arcuate rows; palp represented by a cluster of 
5 sensilla on a slight elevation. (Material studied: numerous larvae 
from South Africa, courtesy of Dr. W. L. Brown.) 

Genus PACHYSIMA Emery 

Pachysima latifrons Emery: Bernard (1951: 1054-1057) de- 
scribed and figured the young (after W. M. Wheeler). 

Genus VITICICOLA Wheeler 

Vitidcola tessmanni (Stitz) : Bernard (1951: 1054) described 
and figured the larva (after W. M. Wheeler). 

SUBFAMILY MYRMICINAE 

Ettershank (1966: 161, 162): "The larvae of the Formicidae 
have not been used to any extent in taxonomic studies, although 
numerous descriptions and figures of scattered genera and species 
occur in the literature. The only wide-scale comparative larval study 



1973] Wheeler f Wheeler Ant Larvae 209 

that has been attempted is the series of papers by G. C. Wheeler 
(later with J. Wheeler), which constitute a fundamental contribu- 
tion to the subject that will be used for a long time." "Reference to 
all the publications by the Wheelers on myrmicine ants are contained 
in a summary article (G. C. and J. Wheeler 1960). In this paper, 
the authors conclude that three characters are of major importance: 
body profile, mandible shape, and setal form. They recognize 22 
body profiles and 30 mandibular shape categories all of which are 
explained and illustrated." 

Genus MESSOR Forel 

Messor capitatus Latreille: Delage (i968a) gave in a table the 
sizes and abundance of larvae throughout the year. She stated that 
only small larvae overwinter. She (ig68b) discussed larval enzymes 
and digestion. 

Genus PHEIDOLE Westwood 

Kempf (1972: 457): Ph. vallifica is the host of the eucharitid 
Orasema costaricensis Wheeler and Wheeler. 

Genus MELISSOTARSUS Emery 

Delage-Darchen (19723) : Hairs few, long, with bifid tips. Crude 
sketch of a larva on p. 219. 

Genus CREMATOGASTER Lund 

Delage-Darchen (ig72b) found only three larval stages in C. 
(Neinatocrema) stadelmanni Mayr. Fig. i hairs enlarged; Fig. 2 
and 3 larvae of various stages in side view; Fig. 4 head in anterior 
view. Pilosity is taxonomically worthless because of extreme vari- 
ation between colonies and even in the same colony. 

Genus MONOMORIUM Mayr 

Cloudsley-Thompson (1962: 179): The calliphorid flies Ben- 
galia peuhi Vil. and B. minor Malloch fed on the larvae of M. 
salomonis (Linnaeus) in the central Sudan. 

Van Pelt and Van Pelt (1972: 978): Larvae of the syrphid 
Microdon baliopterus Loew fed upon the larvae of M. minimum 
(Buckley). 



2IO Psyche [September 

Genus SOLENOPSIS Westwood 

Markin et al. (1972: 1053): Life cycle of Solenopsis invicta 
Buren in an incipient colony: egg 6-8 days, larva 14-15 days, pupa 
20-24 days. 

Genus TETRAMORIUM Mayr 

Tetramorium caespitum (Linnaeus). Donisthorpe (1927: 197) : 
"The larvae were fed with disgorged liquid food as long as they 
were young and gathered together in groups, but when they grew 
older and were separated, the workers fed them with solid sub- 
stances." Many larvae were hung on to the plaster walls of the nest 
by their anchor-tipped hairs. 

LITERATURE CITED 

BERNARD, F. 

1951. Super-famille des Formicoidea. Traite de Zoologic, Tome X, 

Fasc. II: 907-1119, 1258-1263, 1272-1275. 
CLOUDSLEY-THOMPSON, J. L. 

1962. A note on the association between Bengalia spp. (Dipt., Cal- 

liphoridae) and ants in the Sudan. Entomol. Monthly Mag. 98 : 

177-179. 
DELAGE, BERNADETTE. 

1968a. Recherches sur la fourmis moissoneuses du Bassin Aquitain: 

ecologie et biologie. Bull. Biol. 100: 315-367. 
1968b. Recherches sur les fourmis moissoneuses du Bassin Aquitain: 

ethologie. Physiologie de Palimentation. Ann. Sci. Nat., Zool. 

Biol. Anim. (12)10: 197-265. 
1972a. Une fourmi de Cote-d'Ivoire : Melissotarsus titubans Del., n. sp. 

Insectes Sociaux 19: 213-226. 
1972b. Le polymorphisme larvaire chez les fourmis Nematocrema 

d'Afrique. Insectes Sociaux 19: 257-277. 

DONISTHORPE, H. ST. J. K. 

1927. British ants. Geo. Routledge & Sons, London. 436 pp. 

ETTERSHANK, G. 

1966. A generic revision of the world Myrmicinae related to Solenopsis 

and Pheidologeton. Australian J. Zool. 14: 73-171. 
JANZEN, D. H. 

1966. Coevolution of mutualism between ants and acacias in Central 
America. Evolution 20: 249-275. 

1967. Interaction of the bull's-horn acacia (Acacia cornigera L.) with 
an ant inhabitant (Pseudomyrmex ferruginea F. Smith) in eastern 
Mexico. Univ. Kansas Sci. Bull. 47: 315-558. 

KEMPF, W. W. 

1972. A study of some Neotropical ants of genus Pheidole Westwood. I. 
Studia Entomol. 15: 449-464. 



1973] Wheeler fcf Wheeler Ant Larvae 21 1 

MARKIN, G. P., H. L. COLLINS AND J. H. DILLIER. 

1972. Colony founding by queens of the red imported fire ant, Solenop- 

sis invicta. Ann. Entomol. Soc. Amer. 65: 1053-1058. 
SUDD, J. M. 

1967. An introduction to the behavior of ants. St. Martin's Press, New 

York. 200 pp. 
VAN PELT, A. F., AND S. A. VAN PELT. 

1972. Microdon (Diptera: Syrphidae) in nests of Monomorium in 

Texas. Ann. Entomol. Soc. Amer. 65: 977-979. 
WHEELER, G. C., AND JEANETTE WHEELER. 

1956. The ant larvae of the subfamily Pseudomyrmecinae. Ann. Ento- 
mol. Soc. Amer. 49: 374-398. 
1960. Supplementary studies on the larvae of the Myrmicinae. Proc. 

Entomol. Soc. Washington 62: 1-32. 
1964. The ant larvae of the subfamily Cerapachyinae: supplement. 

Proc. Entomol. Soc. Washington 66: 65-71. 
1972. Ant larvae of the subfamily Myrmicinae: second supplement on 

tribes Myrmicini and Pheidolini. J. Georgia Entomol. Soc. 7: 

233-246. 
1973a. The ant larvae of six tribes: second supplement. J. Georgia 

Entomol. Soc. 8: 27-39. 

1973b. Ant larvae of four tribes: second supplement. Psyche 80: 70-82. 
1973c. Ant larvae of the myrmicine tribe Attini: second supplement. 

Entomol. Soc. Washington. (In press.) 
1973d. The ant larvae of the tribes Basicerotini and Dacetini: second 

supplement. Pan-Pacific Entomol. (In press.)